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A Higher Level Classification of All Living

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RESEARCH ARTICLE
A Higher Level Classification of All Living

Organisms
Michael A. Ruggiero1*, Dennis P. Gordon2, Thomas M. Orrell1, Nicolas Bailly3,
Thierry Bourgoin4, Richard C. Brusca5, Thomas Cavalier-Smith6, Michael D. Guiry7, Paul
M. Kirk8
1 Integrated Taxonomic Information System, National Museum of Natural History, Smithsonian Institution,
Washington, District of Columbia, United States of America, 2 National Institute of Water & Atmospheric
Research, Wellington, New Zealand, 3 WorldFishFIN, Los Baos, Philippines, 4 Institut Systmatique,
Evolution, Biodiversit (ISYEB), UMR 7205 MNHN-CNRS-UPMC-EPHE, Sorbonne Universits, Museum
National d'Histoire Naturelle, 57, rue Cuvier, CP 50, F-75005, Paris, France, 5 Department of Ecology &
Evolutionary Biology, University of Arizona, Tucson, Arizona, United States of America, 6 Department of
Zoology, University of Oxford, Oxford, United Kingdom, 7 The AlgaeBase Foundation & Irish Seaweed
Research Group, Ryan Institute, National University of Ireland, Galway, Ireland, 8 Mycology Section, Royal
Botanic Gardens, Kew, London, United Kingdom
* ruggierm@si.edu
Abstract
OPEN ACCESS
Citation: Ruggiero MA, Gordon DP, Orrell TM, Bailly
N, Bourgoin T, Brusca RC, et al. (2015) A Higher
Level Classification of All Living Organisms. PLoS
ONE 10(4): e0119248. doi:10.1371/journal.
pone.0119248
Academic Editor: Erik V. Thuesen, The Evergreen
State College, UNITED STATES
Received: July 8, 2014
Accepted: January 25, 2015
Published: April 29, 2015
Copyright: This is an open access article, free of all
copyright, and may be freely reproduced, distributed,
transmitted, modified, built upon, or otherwise used
by anyone for any lawful purpose. The work is made
available under the Creative Commons CC0 public
domain dedication.
We present a consensus classification of life to embrace the more than 1.6 million species
already provided by more than 3,000 taxonomists expert opinions in a unified and coherent,
hierarchically ranked system known as the Catalogue of Life (CoL). The intent of this collaborative effort is to provide a hierarchical classification serving not only the needs of the
CoLs database providers but also the diverse public-domain user community, most of
whom are familiar with the Linnaean conceptual system of ordering taxon relationships.
This classification is neither phylogenetic nor evolutionary but instead represents a consensus view that accommodates taxonomic choices and practical compromises among diverse
expert opinions, public usages, and conflicting evidence about the boundaries between
taxa and the ranks of major taxa, including kingdoms. Certain key issues, some not fully resolved, are addressed in particular. Beyond its immediate use as a management tool for the
CoL and ITIS (Integrated Taxonomic Information System), it is immediately valuable as a
reference for taxonomic and biodiversity research, as a tool for societal communication,
and as a classificatory backbone for biodiversity databases, museum collections, libraries,
and textbooks. Such a modern comprehensive hierarchy has not previously existed at this
level of specificity.
Data Availability Statement: All relevant data are

within the paper and its Supporting Information files.
Funding: The authors have no funding or support to
report.
Competing Interests: The authors have declared
that no competing interests exist.
Introduction
Biological classification (taxonomy) aims to simplify and order the immense diversity of life
into coherent units called taxa that have widely accepted names and whose members share important properties. It synthesizes information concerning a great variety of characters (e.g.,
PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015
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A Higher Level Classification of All Living Organisms
morphological; molecular: genes, metagenome, and metabolome; etho-ecological). There is

currently no consensus among the world's taxonomists concerning which classification scheme
to use for the overall hierarchy of life, in part because of the confusion resulting from Hennig's
[1] redefinition of previous terminology of classification, which has not been universally accepted; the separate goals of cladification and classification [2]; and conflicting or unresolved
evidence for phylogenetic relationships. The continuing advances in the use of specialized analytical tools from many different fields and their resulting conclusions and assumptions require
regular updates as advances in knowledge are made.
Biological classification can integrate diverse, character-based data in a phylogenetic framework, which allows a broad user community to utilize the disparate knowledge of shared biological properties of taxa. Phylogeny is, therefore, the basis for these biological classifications
but there is still strong debate over their accounting for evolutionary divergence or information
content other than the branching pattern [3]. Accordingly, classifications have often been labeled either phylogenetic or evolutionary, depending mainly upon whether or not they reject
paraphyletic groups [3, 4].
While the type of classification to be used to support further exploration and analysis of any
biological scenario may be important, it is not the subject of this paper. The proposed classification does not address detailed phylogenetic questions and, while hierarchical and reflective
of phylogeny, is not itself a phylogenetic tree. The aim of this classification is to be a pragmatic
means of managing the ever-increasing knowledge of the diversity of life, its relationships,
characteristics, and properties. Indeed, the past two decades have witnessed an explosion in
biodiversity research and informatics, emphasizing the need for a quality list of accepted scientific names of the more than 1.9 million described living species [5] and for greater consensus
on how to classify them at higher taxonomic ranks. Since 2001, Species 2000 and the Integrated
Taxonomic Information System (ITIS) have worked with their respective contributors to complete a comprehensive species list, called the Catalogue of Life (CoL). The CoL Annual Checklist (http://www.catalogueoflife.org/annual-checklist/2014/) already contains more than 1.6
million valid or accepted species names provided by more than 140 taxonomic databases involving more than 3,000 taxonomists [6]. More than 82% of the global species databases are
provided at the rank of class or below (includes 1.3 million species), and more than 63% are
provided at the rank of order or below (includes 1.0 million species). Owing to the heterogeneity in higher level classification among the contributed databases, the CoL managers sought a
practical and coherent hierarchical classification that could serve as a framework for data integration. Here we explain the rationale behind the consensus higher level classification that we
propose for CoL use.
Our goal, therefore, is to provide a hierarchical classification for the CoL and its contributors that (a) is ranked to encompass ordinal-level taxa to facilitate a seamless import of contributing databases; (b) serves the needs of the diverse public-domain user community, most of
whom are familiar with the Linnaean conceptual system of ordering taxon relationships; and
(c) is likely to be more or less stable for the next five years. Such a modern comprehensive hierarchy did not previously exist at this level of specificity. In this sense it summarizes overarching
aspects of the tree of life, including both paraphyletic and monophyletic groups, both being important in facilitating meaningful communication among scientists and between the scientific
community and society.
The most recent higher level classification to this level was published more than 30 years
ago, before the advent of modern molecular analysis [7]. Beyond the immediate use for CoL,
the hierarchy is valuable as a reference for taxonomic and biodiversity research, as a tool for societal communication, and as a stable classificatory backbone for biodiversity databases, museum collections, libraries, and textbooks, to name a few applications.
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Approach
When Linnaeus introduced his novel system of nature in the mid-18th century, he recognized
three kingdoms of nature: Regnum Vegetabile (plants), Regnum Animale (animals), and Regnum Lapideum (minerals) that has long since been abandoned. However, as is evident from
the title of his work, he introduced lower level taxonomic categories (named class, order,
genus, and species), each successively nested within higher ranked categories. Linnaeus' system
has proven to be robust for more than 250 years (see the comprehensive discussion and suggestions for dealing with potential conflicts in Vences et al. [8]). In modern-day classifications, the
starting point for botanical names is Linnaeus Species Plantarum [9] and for zoological names
it is the tenth edition of the Systema Naturae [10]. Since Linnaeus, the expansion of knowledge
and the increase in the number of described species has required an expansion of the number
of hierarchical levels (ranks) within the system. The categories of family and phylum (or division) were introduced in the early 19th century and many intermediate categories have been
added since. There is currently little agreement about the general names for categories above
that of kingdom; here we use superkingdom rather than empire or domain. In addition, there
are three separate codes that govern the assignment and use of scientific names, each with different requirements and terminology and consequences for their classifications. For algae,
fungi, and plants (ICN: International Code of Nomenclature for algae, fungi, and plants), the
principle of priority does not apply above rank of family; for animals (ICZN: International
Code of Zoological Nomenclature), priority does not apply above the family-group ranks; and
for prokaryotes other than Cyanobacteria (ICNB: International Code of Nomenclature of Bacteria), only the categories ranked as class and below are covered by the code. A recent paper
by the International Committee on Bionomenclature compares terminology among six
current nomenclatural codes and makes recommendations for their use in improving communication [11].
In 2005, on behalf of the International Society of Protistologists, Adl et al. [12] presented a
nested eukaryote-only cladification that used the names of six supergroupsAmoebozoa,
Opisthokonta, Rhizaria, Excavata, Chromalveolata, and Archaeplastida (= Plantae) [1317]
as the highest ranked eukaryote groups. Their schema was updated in 2012 [18], with Rhizaria
and Chromalveolata replaced by SAR plus four small hacrobian groups. Although these taxa
are nested, and ranked by a bulleted system, Adl et al. avoided the use of Linnaean higher category names (phylum, class, order, family) that would have more usefully denoted rank. Insofar as the nested groups comprise a mix of taxon names based on priority (i.e., according to the
year of introduction of the name), many individual genera as well as traditional taxon names
(family through class) end up having the same rank in the Adl et al. hierarchy, while at the
same time having different suffixes or none at all. The ranks assigned therein often seem to
reflect our present partial ignorance of relationships more than careful assessment of relative
phenotypic disparity as in Linnaean taxonomy. This is very confusing when these group
names (genus to kingdom) are used in isolation without regard to phylogenetic relativity.
Two of the great benefits of Linnaean-ranked categories and their standardized suffixes are
that they instantly relativize taxa that are otherwise unknown to the non-specialist and also
indicate the relative degree of phenotypic distinctiveness amongst groups. The overarching
higher level classification used by the CoL, therefore, uses the standard formal categories, as
it is intended to be simultaneously pragmatic and informative of both evolutionary relatedness and relative phylogenetic subordination. A classification should be biologically wellgrounded and widely useful. In its simplicity, it provides less detail about relationships than a
complete phylogeny but is still congruent with it [19]. Our classification is not intended to
compete with a cladification such as Adl et al.sboth are valid ways of ordering the living
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worldbut we would argue that theirs is less comprehensible to many in the public-domain
user communities.
These actual complexities of phylogenetic history emphasize that classification is a practical
human enterprise where compromises must be made [20]. We have therefore named only
groups generally considered to have had a monophyletic origin, even though some of them
may be paraphyletic (i.e., do not include all descendants of their last common ancestor) and
others, e.g., Euglenozoa, Rhizaria, Cercozoa, include subgroups (such as Euglenophyceae,
Chlorarachnea, and Paulinella) that evolved by the symbiogenetic merger of two fundamentally different lineages [21], while others have had infusions of genes from elsewhere [22] and
therefore do not conform to any purely formal definition of monophyly. We have not adopted
the view that one should never accept paraphyletic groups in a classification but rather have
evaluated each case of paraphyly on its practicability and usage. In some cases (e.g., classical
bryophytes) we accepted the splitting of paraphyletic taxa into holophyletic groups (groups
with a monophyletic origin that also include all descendants of their last common ancestor, i.e.,
clades). In others we retained ancestral (paraphyletic) taxa when it seemed beneficial to do so
(e.g., Prokaryota, Protozoa, Crustacea, Sarcopterygii, Reptilia). For practical purposes we treat
Proteobacteria and Cyanobacteria as holophyletic phyla even though both exclude their mitochondrial and chloroplast descendants, neither of which is now a bacterium but an evolutionarily chimaeric cell organelle. We have conservatively retained several groups where evidence
for paraphyly or holophyly is contradictory, such as Archaea (Archaebacteria).
A panel of experts representing the major taxonomic disciplines was convened to review, revise, and update the existing incomplete CoL hierarchy. These authors consulted more than
200 sources (see S1 Appendix), most of which were from recent taxonomic publications and
websites. The product is a current and practical classification that meets the panels established
goal. In achieving a consensus, the panel was required to make some compromises that may require future revision as the related issues are resolved. While all of these individuals made contributions to the hierarchy, not all necessarily endorse every aspect of it. The CoL classification
will undergo review and revision at five-year intervals to consider changes as necessary.
Results and Discussion

We are proposing a two-superkingdom (Prokaryota and Eukaryota), seven-kingdom classification that is a practical extension of Cavalier-Smiths six-kingdom schema [19]; the latter has
been used, for example, in the compendious checklist of marine biota of Chinese seas [23] and
in the first comprehensive national inventory of biodiversity for New Zealand [2426]. For
each of these kingdoms we had to exercise our taxonomic judgment and reach a practical compromise among diverse opinions and usages and conflicting evidence about certain phylogenetic questions important for defining the boundaries between and ranks of major taxa,
including kingdoms. Our schema includes: the prokaryotic kingdoms Archaea (Archaebacteria) and Bacteria (Eubacteria), and the eukaryotic kingdoms Protozoa, Chromista, Fungi,
Plantae, and Animalia. We have retained 14 ranks from superkingdom to order (Table 1). Several key taxonomic issues, some not fully resolved, are discussed below.
Prokaryota
The higher classification of prokaryotes is still somewhat unsettled. Woese and Fox [27] treated
Archaebacteria (Archaea) and Eubacteria (Bacteria) as separate kingdoms. Margulis and
Schwartz [28] recognized the superkingdom Prokarya, containing one kingdom Bacteria that
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Table 1. List of ranks used in the hierarchy with the number of taxa per rank.
Rank
Number of Taxa
Superkingdom
Kingdom
Subkingdom
11
Infrakingdom
Superphylum
Phylum
96
Subphylum
60
Infraphylum
Superclass
12
Class
351
Subclass
145
Infraclass
23
Superorder
52
Order
1,467
Main ranks are in bold type; unnamed taxa are not counted.
doi:10.1371/journal.pone.0119248.t001
included a subkingdom Archaea; Cavalier-Smith also treated Archaebacteria and Eubacteria as

prokaryote subkingdoms [19, 29]. Commonly used sources of prokaryote names, such as the
List of Prokaryotic Names with Standing in Nomenclature (LPSN) [30] and the Taxonomic
Outline of Bacteria and Archaea (TOBA) [31] treat Bacteria and Archaea as separate domains
but are silent about the category of kingdom. While these sources list the names of phyla in
common use as a service to the user, they are not validly published under the ICNB. We have
not placed phylum names in quotation marks as they have but we have so designated a few
prokaryote names at lower ranks that are in common use but not (or not yet) valid. As no prokaryote names above the ranks of class are covered by ICNB rules, there is no official higher
classification of prokaryotes [32] and any attempt at such is necessarily difficult. We have chosen to adopt the classification in current use by the Catalogue of Life. It is derived from the
TOBA and recognizes Bacteria and Archaea as equivalent in rank to the eukaryote kingdoms.
We treat them as de facto kingdoms until there is a better resolution of their status. The number of negibacterial phyla currently recognized [30] is probably excessive compared with eukaryotes and mainly reflects uncertainty about the true relationships of many small phyla,
probably exaggerating the significance of their biological disparity. Greater use of multigene
trees rather than over reliance on rRNA gene trees alone may eventually allow further
simplification by grouping them into fewer phyla, possibly only about half the present
number [28].
Protozoa and Chromista

Unicellular eukaryotes, usually called protists, comprise a polyphyletic group of eukaryotes
that do not undergo tissue formation through the process of embryological layering. They include ancestrally unicellular eukaryotes directly descended from bacteria by the origin of the
nucleus, endomembrane, cytoskeleton, and mitochondria. Assigning them to separate kingdoms was historically difficult when only light microscopy was available but is now
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considerably facilitated because of advances in electron microscopy and gene sequencing. Formerly, the unicellular amoeboid group Myxozoa with multicellular spores was included in Protozoa but these protists are now firmly within the animal kingdom, having been proven to be
greatly simplified parasitic animals. Yeasts are unicellular fungi that evolved polyphyletically
from multicellular filamentous ancestors and are assigned to one of three higher fungal phyla.
Microsporidia are highly reduced intracellular parasites traditionally considered to be Protozoa, but they have been known for two decades to be related to Fungi. At one time it was
thought microsporidia had evolved from Fungi and therefore were placed in that kingdom [19,
33]. For several years multigene trees were contradictory about whether microsporidia
branched within or diverged from Fungi. The latest evidence is that they are most closely related to rozellids [34], which also have been treated either as Fungi or Protozoa. If this recent phylogeny [34] is correct, both should be in the same kingdom. Here we take the view that the best
demarcation between Protozoa and Fungi lies immediately before the origin of the chitinous
wall around vegetative fungal cells and associated loss of phagotrophy [33]. We therefore include microsporidia and rozellids in Protozoa (vegetatively wall-less, typically phagotrophs)
not Fungi (vegetatively walled osmotrophs).
For decades, taxonomists have debated the boundary between Protozoa and Plantae. We accept the view that it should be placed just prior to the evolutionary origin of chloroplasts and
that Plantae should comprise all eukaryotes with plastids directly descending from the initially
enslaved cyanobacterium, i.e., Viridiplantae (green plants), Rhodophyta (red algae), and Glaucophyta (glaucophyte algae), but exclude those like chromists that got their chloroplasts from
plants secondarily by subsequent eukaryote-to-eukaryote lateral transfers. Therefore, all green
algae are included in Viridiplantae and Plantae and are excluded from Protozoa. The only photosynthetic Protozoa are Euglenophyceae, which obtained their chloroplasts subsequently from
an enslaved green alga [21].
The boundary between Protozoa and Chromista has been more controversial. Chromista
was established to include all chromophyte algae (those with chlorophyll c, not b) considered
to have evolved by symbiogenetic enslavement of another eukaryote (a red alga) as well as all
heterotrophic protists descended from them by loss of photosynthesis or entire plastids [35].
With phylogenetic advances it has become clearer that alveolates (once considered Protozoa)
are related to chromistan heterokont algae (and related heterotrophic heterokonts) and more
distantly to Rhizaria, the three together forming the major group Harosa (equivalent to SAR).
Consequently, Chromista has been greatly expanded to include all Harosa as well as other former protozoa that turned out to be related to haptophytes or cryptophytes. Chromista now includes many groups once treated as Protozoa [19], an expansion followed here. In multigene
trees, this expansion is the most difficult part of the entire eukaryote tree to resolve. They sometimes show one or both of Plantae and Chromista as a clade but often their major subgroups
are intermingled in contradictory ways [36, 37]. This may be a consequence of the eukaryoteeukaryote chimaeric history of chromists that acquired some genes from red algae or of the
very rapid basal radiation of the robust corticate clade (i.e. Plantae plus Chromista). Because of
this, some question whether Chromista represents a clade, yet trees are still too poorly resolved
to eliminate the likelihood from cell evolutionary considerations that Chromista and Plantae
are genuinely distinct sister clades. Evidence that Harosa is a clade is very strong. Evidence
that Haptista plus Cryptista are a clade Hacrobia is strong on some trees but questioned by
others [37].
Protozoa, like Prokaryota, is certainly a paraphyletic taxon [38]; Animalia, Fungi, Plantae,
and Chromista all evolved from it. In our hierarchy Protozoa comprises seven phyla, of which
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four are probably clades and three paraphyletic. We do not consider it useful in a general classification to subdivide the paraphyletic phyla into numerous smaller ones, often with only a
handful of species that most have never heard of, even though a few specialists might favor that
despite their constituent subgroups not differing radically in cell structure. For both Protozoa
and Chromista we have favored large groups with shared body plans, analogous to extremely
diverse animal phyla like Chordata and Arthropoda. The higher proportion of ancestral (paraphyletic) phyla in Protozoa compared with terminal groups like animals and plants is unsurprising because they were the first eukaryotes and they diverged early on but with many fewer
associated major changes in body plan than occurred during the much later radiation of bilateral animals. Distinct early diverging protozoan clades can be remarkably similar morphologically and biologically [39].
Fungi
As stated earlier, we take the view that the best demarcation between Protozoa and Fungi lies
immediately before the origin of the chitinous wall around vegetative fungal cells and associated loss of phagotrophy. We use an updated version of the higher classification presented in the
10th Edition of the Dictionary of Fungi [40]. The evolutionarily convergent Oomycetes such as
the serious pest Phytophthora, formerly treated as Fungi, belong instead in phylum Pseudofungi of the heterokont Chromista.
Plantae
As with the other kingdoms, Plantae is classified in a variety of ways. Margulis and Schwartz
[28] restricted Plantae to land plants (embryophytes or higher plants) and popularized the use
of kingdom Protoctista to include lower plants (green, red, and glaucophyte algae) and lower
Fungi as well as chromists with classical protozoa. Many now consider such a kingdom too
broad and heterogeneous and the associated separation of lower and higher plants in different
kingdoms to be undesirable. Now taxonomists almost universally classify lower and higher
plants together in the single kingdom Plantae and lower and higher fungi within the single
kingdom Fungi. We have adopted this delimitation of Plantae here [19, 35] (for which Archaeplastida [12, 18] is a less familiar recent synonym). The structure of plastid genomes and the
derived chloroplast protein-import machinery support a single origin of glaucophytes, red
algae, green algae, and embryophytes (land plants). The ancestral embryophyte is thought to
have originated from relatives of the Charales (stoneworts) or Coleochaetales (Charophyta).
Jeffrey [41] first grouped charophytes and embryophytes as a clade Streptophyta, which was
later validated as a superphylum [42] and reduced to phylum by Bremer [43].
Chase and Reveal [44] published a phylogenetic classification of land plants, reasoning that
If the major clades of green algae are recognized as classes, then all land plants, the embryophytes, should be included in a single class, here recognized as Equisetopsida. This argument,
however, overemphasizes cladistic level compared with phenotypic disparity, and is contrary to
traditional assignment of phylum (or division) status to the main bryophyte, pteridophyte
and seed-plant subgroups. This latter treatment was exemplified in the 2008 Annual Checklist
of the CoL, which listed three bryophyte phyla, four pteridophyte phyla, and five seed-plant
phyla, reflecting the arrangement found in many university textbooks of the late 20th century
and in Margulis and Schwartzs Five Kingdoms [28]. Here we recognize four embryophyte
phylathree of bryophytes (liverworts, hornworts, and mosses) and a single phylum Tracheophyta for vascular plantswith all species characterized by a diploid phase having xylem
and phloem. Bryophyte specialists tend to treat each of the three major bryophyte groups
as phylaMarchantiophyta, Anthocerotophyta, Bryophyta [45, 46]. We have chosen a
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conservative approach to the higher classification of plants, largely consistent with Mabberley
[47] for the embryophyte ranks above class, while using Chase and Reveal [44] and Stevens
[48] for the lower ranks.
Animalia (Metazoa)
The numbers of phyla and classes with extant species in kingdom Animalia differ according to
molecular and morphological partitioning in phylogenies [4959] as well as the preferred treatments of specialists of particular traditional phyla and where to draw the line between related
taxa and how to rank themthe ranking of phylum versus subphylum is sometimes rather
subjective. Based on the contributions of taxonomic experts to an outline of higher level classification and survey of taxonomic richness [60, 61], as many as 39 animal phyla might be recognized (more, if Porifera were abandoned as a phylum and constituent major clades given
higher rank [62]). Below we discuss some issues encountered in arriving at decisions for our
proposed classification, which accepts 34 animal phyla.
(1) Poriferaone phylum or three? Nielsen [62] argued that The three apparently
monophyletic sponge groups Silicea, Calcarea, and Homoscleromorpha do not constitute a
monophyletic group, and the phylum Porifera thus has to be abandoned. More recent studies alternatively support paraphyly [63] or holophyly [58, 64] of sponges. Until the issue is resolved, we will follow the Porifera community [6567] in retaining one phylum Porifera with
four classes.
(2) Status of Myxozoa. Recent work on the vermiform myxozoan Buddenbrockia has
demonstrated conclusively that myxozoans are extremely simplified Cnidaria, possibly Medusozoa [68, 69]. We classify Myxozoa as a subphylum of Phylum Cnidaria.
(3) Flatwormsmonophyletic or not? In 1995, Nielsen [70] wrote The delimitation of
the phylum [Platyhelminthes] is not much in question, but recent molecular analyses, combined with a careful reconsideration of morphology and anatomy, have confused the classification of Platyhelminthes, affecting particularly Acoela, Xenoturbella, and Nemertodermatida.
Egger et al. [71] reviewed the evidence, noting the contrast between morphological and phylogenomic data. Whereas the stem-cell system and the mode of replacing epidermal cells unite
both Acoela and Rhabditophora and are not found in any other bilaterian lineage, phylogenomic data support a separation of these two groups, a conclusion reached by Philippe et al.
[72] based on mitochondrial genes, a phylogenomic data set of 38,330 amino-acid positions,
and miRNA complements. We follow Philippe et al. [72] and Tyler and Schilling [73] in uniting Acoela, Xenoturbella, and Nemertodermatida as the deuterostome phylum Xenacoelomorpha. The remaining internal classification of Platyhelminthes is also somewhat problematic.
We propose a classification that is based in part on Riutort et al. [74] and Tyler [75].
(4) Phylum Gnathifera or phyla Acanthocephala, Gnathostomulida, Micrognathozoa,
and Rotifera? Until recently, all four of these groups were commonly treated as separate
phyla [28, 61, 7680]. However, numerous recent molecular and morphological analyses nest
Acanthocephala within Rotifera [8186]. A syncytial epidermis links rotifers, Seison and
Acanthocephala; Ahlrichs [87, 88] proposed Syndermata for this clade. As revealed by transmission electron microscopy [89] and scanning electron microscopy [90], the jaw apparatus of
gnathostomulids and rotifers is remarkably similar. That of Seison is less obviously homologous [91] and the Seisonidea may have diverged from rotifers at an early stage of their evolution. On the other hand, Seison has similar sperm to acanthocephalans and the epidermis of
both groups contains bundles of filaments. Limnognathia maerski, representing a new category
of organism (Micrognathozoa) from cold fresh waters in Greenland and the Crozet Islands [92,
93], has a remarkable jaw apparatus (the most complicated known among invertebrates) with
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clear homologies, in both the jaw elements and musculature, with the trophi in Rotifera and
the jaws in Gnathostomulida. The jaw apparatus and musculature, as well as molecular analyses, unite these taxa as a clade known as Gnathifera (see [86, 92]). In the analysis by Giribet
et al. [94], the issue remained unresolved, as Micrognathozoa appeared independent of
Gnathostomulida and Rotifera, with unclear affiliation. Edgecombe et al. [59] and Nielsen [95]
retain phylum status for Gnathostomulida, Micrognathozoa, and Rotifera but not Acanthocephala. We treat each of the major gnathiferan groups as a phylum, including Acanthocephala,
following Monks and Richardson [79], though some of us think that the number of gnathiferan
phyla ought to be substantially reduced when their phylogeny, including ingroup relationships
of Rotifera sensu lato, is more firmly established.
(5) The scalidophoran phyla Adrianov and Malakhov [96] erected phylum Cephalorhyncha for Kinorhyncha, Loricifera, Priapula, and Nematomorpha. The first three of these
phyla have in common an eversible snout (introvert) with scalid spines and inner and outer retractor muscles, a similar excretory filter (protonephridium), and similar sense organs, providing strong justification for uniting them in a single clade, the Scalidophora [97]. There is also
molecular support, though not unanimity, for a clade of Kinorhyncha, Loricifera, and Priapula,
known as Scalidophora. On the other hand, Kinorhyncha has internal and external body segmentation lacking in the other groups. Neuhaus and Higgins [98] noted that conflicting evidence exists for every one of the possible sister-group relationships among these phyla and
prefer to keep them separate in a superphylum Scalidophora (which is preferred over Cephalorhyncha, the latter name originally including the Nematomorpha). We recommend separate
scalidophoran phyla, though the number might be greatly reduced when the phylogeny
becomes clearer.
(6) The chordate subphyla Cephalochordata and Urochordata Some sequence analyses
have questioned the monophyly of Chordata [99, 100]. Nielsen [95] maintains Urochordata
(or Tunicata) and Cephalochordata as separate phyla, whereas the group Urochordata is closer
to Vertebrata (craniates), in a clade Olfactores, than Cephalochordata. We retain all three
groups as traditional chordate subphyla.
Many users of classifications would prefer a stable, unchanging system. Yet classifications
are syntheses of biological knowledge, particularly contemporary phylogenetic understanding
of taxa, that must be regularly updated in accord with new scientific discoveries. Taxonomy
must therefore navigate between the dual perils of ignoring important advances and making
premature or unnecessary changes. We seek stability in nomenclature at the species level but at
higher levels the concepts and compositions of major taxa, and therefore the scope of wellknown names, must inevitably shift as new organisms are discovered and evolutionary affinities are better understood. The fact that we have been able to agree on a practical unified classification shows that taxonomists can broadly agree, despite the diverse experiences, viewpoints,
and to some extent, differing philosophies of classification represented on our panel. The present classification (as, indeed, all classifications) should be regarded as interim, and it will inevitably change in certain respects, some hinted at above. However, we suspect that the recent
torrent of radical re-evaluations (resulting especially from the application of DNA sequencing
and other new techniques) may lessen as time passes. We hope that this unusually comprehensive classification will be widely useful and provide a sound basis for further improvement.
A complete proposed classification from superkingdom to order is provided in Table 2 and
is available for download at <http://www.catalogueoflife.org/col/>. Below the rank of infrakingdom, we have followed the convention used in the Catalogue of Life and listed taxon
names alphabetically. This allows easier searching by those not familiar with the phylogenies of
9 / 60
Table 2. Proposed hierarchical classification from superkingdom to order.

SUPERKINGDOM PROKARYOTA
KINGDOM ARCHAEA [= ARCHAEBACTERIA]
Phylum Crenarchaeota
Class Aigarchaeota
Order N.N. ("Ca. Caldiarchaeum")
Class Korarchaeota
Order N.N. ("Ca. Korarchaeum")
Class Thaumarchaeota
Order Cenarchaeales
Class Thermoprotei [= Crenarchaeota]
Order Acidilobales
Order Desulfurococcales
Order Fervidicoccales
Order Sulfolobales
Order Thermoproteales
Phylum Euryarchaeota
Class Archaeoglobi
Order Archaeoglobales
Class Halobacteria
Order Halobacteriales
Class Methanobacteria
Order Methanobacteriales
Class Methanococci
Order Methanococcales
Class Methanomicrobia
Order N.N. (Methanocalculus)
Order Methanocellales
Order Methanomicrobiales
Order Methanosarcinales
Class Methanopyri
Order Methanopyrales
Class Nanohaloarchaea
Order N.N. (e.g., "Ca. Nanosalinarum")
Class Thermococci
Order Thermococcales
Class Thermoplasmata
Order Thermoplasmatales
KINGDOM BACTERIA [= EUBACTERIA]
SUBKINGDOM NEGIBACTERIA
Phylum Acidobacteria
Class N.N. (Bryobacter)
Class Acidobacteria
Order Acidobacteriales
Class Holophagae
Order Acanthopleuribacterales
Order Holophagales
Phylum Aquicae
Class Aquicae
Order Aquicales
Phylum Armatimonadetes
(Continued)
10 / 60
Table 2. (Continued)
Class Armatimonadia
Order Armatimonadales
Class Chthonomonadetes
Order Chthonomonadales
Class Fimbriimonadia
Order Fimbriimonadales
Phylum Bacteroidetes
Class Bacteroidia
Order Bacteroidales
Class Cytophagia
Order Cytophagales
Class Flavobacteria
Order Flavobacteriales
Class Sphingobacteriia
Order Sphingobacteriales
Phylum Caldiserica
Class Caldisericia
Order Caldisericales
Phylum Chlamydiae
Class Chlamydiae
Order Chlamydiales
Phylum Chlorobi
Class Chlorobia
Order Chlorobiales
Class Ignavibacteria
Order Ignavibacteriales
Phylum Chrysiogenetes
Class Chrysiogenetes
Order Chrysiogenales
Phylum Cyanobacteria [= Cyanophyta]
Class Cyanophyceae [= Phycobacteria]
Order Chroococcales
Order Nostocales
Order Oscillatoriales
Order Pseudoanabaeniales
Order Synechococcales
Class Gloeobacteria [= Gloeobacterophyceae]
Order Gloeobacterales
Phylum Deferribacteres
Class Deferribacteres
Order Deferribacterales
Phylum Deinococcus-Thermus [= Hadobacteria]
Class Deinococci
Order Deinococcales
Order Thermales
Phylum Dictyoglomi
Class Dictyoglomia
Order Dictyoglomales
Phylum Elusimicrobia
Class Elusimicrobia
(Continued)
11 / 60
Order Elusimicrobiales
Phylum Fibrobacteres
Class Fibrobacteria
Order Fibrobacterales
Phylum Fusobacteria
Class Fusobacteriia
Order Fusobacteriales
Phylum Gemmatimonadetes
Class Gemmatimonadetes
Order Gemmatimonadales
Phylum Lentisphaerae
Class Lentisphaeria
Order Lentisphaerales
Order Victivallales
Class Oligosphaeria
Order Oligosphaerales
Phylum Nitrospira
Class "Nitrospira"
Order "Nitrospirales"
Phylum Planctomycetes
Class Phycisphaerae
Order Phycisphaerales
Class Planctomycea
Order Planctomycetales
Phylum Proteobacteria
Class Alphaproteobacteria
Order N.N. (e.g., Breoghania)
Order Caulobacterales
Order Kiloniellales
Order Kordiimonadales
Order Magnetococcales
Order Parvularculales
Order Rhizobiales
Order Rhodobacterales
Order Rhodospirillales
Order Rickettsiales
Order Sneathiellales
Order Sphingomonadales
Class Betaproteobacteria
Order N.N. (Chitinivorax)
Order Burkholderiales
Order Hydrogenophilales
Order Methylophilales
Order Neisseriales
Order Nitrosomonadales
Order "Procabacteriales"
Order Rhodocyclales
Class Deltaproteobacteria
Order N.N. (e.g., Deferrisoma)
Order Bdellovibrionales
(Continued)
12 / 60
Order Desulfarculales
Order Desulfobacterales
Order Desulfovibrionales
Order Desulfurellales
Order Desulfuromonadales
Order Myxococcales
Order Syntrophobacterales
Class Epsilonproteobacteria
Order Campylobacterales
Order Nautiliales
Class Gammaproteobacteria
Order N.N. (e.g., Alkalimonas)
Order Acidithiobacillales
Order Aeromonadales
Order Alteromonadales
Order Cardiobacteriales
Order Chromatiales
Order Enterobacteriales
Order Legionellales
Order Methylococcales
Order Oceanospirillales
Order Orbales
Order Pasteurellales
Order Pseudomonadales
Order Salinisphaerales
Order Thiotrichales
Order Vibrionales
Order Xanthomonadales
Class Zetaproteobacteria
Order Mariprofundales
Phylum Spirochaetae
Class
"Spirochaetes"
Order Spirochaetales
Phylum Synergistetes
Class Synergistia
Order Synergistales
Phylum Thermodesulfobacteria
Class Thermodesulfobacteria
Order Thermodesulfobacteriales
Phylum Thermotogae
Class Thermotogae
Order Thermotogales
Phylum Verrucomicrobia
Class Opitutae
Order Opitutales
Order Puniceicoccales
Class Verrucomicrobiae
Order Verrucomicrobiales
SUBKINGDOM POSIBACTERIA
(Continued)
13 / 60
Phylum Actinobacteria
Class Actinobacteria
Order Acidimicrobiales
Order Actinomycetales
Order Bidobacteriales
Order Coriobacteriales
Order Euzebyales
Order Gaiellales
Order Nitriliruptorales
Order Rubrobacterales
Order Solirubrobacterales
Order Thermoleophilales
Phylum Chloroexi [= Chlorobacteria]
Class Anaerolineae
Order Anaerolineales
Class Caldilineae
Order Caldilineales
Class Chloroexia
Order Chloroexales
Order Herpetosiphonales
Class Dehalococcoidia
Order Dehalococcoidales
Class Ktedonobacteria
Order Ktedonobacterales
Order Thermogemmatisporales
Class Thermomicrobia
Order Sphaerobacterales
Order Thermomicrobiales
Phylum Firmicutes
Class Bacilli
Order Bacillales
Order Lactobacillales
Class Clostridia
Order Clostridiales
Order Halanaerobiales
Order Natranaerobiales
Order Thermoanaerobacterales
Class Erysipelotrichia
Order Erysipelotrichales
Class Negativicutes
Order Selenomonadales
Class Thermolithobacteria
Order Thermolithobacterales
Phylum Tenericutes
Class Mollicutes
Order Acholeplasmatales
Order Anaeroplasmatales
Order Entomoplasmatales
Order Haloplasmatales
Order Mycoplasmatales
(Continued)
14 / 60
SUPERKINGDOM EUKARYOTA
KINGDOM PROTOZOA
SUBKINGDOM EOZOA
INFRAKINGDOM EUGLENOZOA
Phylum Euglenozoa
Subphylum N.N.
Class Diplonemea
Order Diplonemida
Class Kinetoplastea
Order Bodonida
Order Prokinetoplastida
Order Trypanosomatida
Subphylum Euglenoida
Class N.N.
Order Petalomonadida
Order Ploeotiida
Class Euglenophyceae
Order Euglenida
Order Eutreptiida
Class Peranemea
Order Heteronemida
Order Peranemida
Order Rhabdomonadida
Subphylum Symbiontida
Class Postgaardea
Order Postgaardida
INFRAKINGDOM EXCAVATA
Phylum Loukozoa
Subphylum Eolouka
Class Jakobea
Order Jakobida
Class Tsukubea
Order Tsukubamonadida
Subphylum Neolouka
Class Malawimonadea
Order Malawimonadida
Phylum Metamonada
Class Anaeromonadea
Order Oxymonadida
Order Trimastigida
Class Carpomonadea
Order Carpediemonadida
Order Chilomastigida
Order Dysnectida
Class Eopharyngea
Order Diplomonadida
Order Retortamonadida
Class Trichomonadea
Order Cristamonadida
Order Spirotrichonymphida
(Continued)
15 / 60
Order Trichomonadida
Order Tritrichomonadida
Class Trichonymphea
Order Lophomonadida
Order Trichonymphida
Phylum Percolozoa
Subphylum Pharyngomonada
Class Pharyngomonadea
Order Pharyngomonadida
Subphylum Tetramitia
Class Heterolobosea
Order Acrasida
Order Schizopyrenida
Class Lyromonadea
Order Lyromonadida
Class Percolatea
Order Percolomonadida
Order Pseudociliatida
SUBKINGDOM SARCOMASTIGOTA
Phylum Amoebozoa
Subphylum Conosa
Class Archamoebea
Order Mastigamoebida
Order Pelobiontida
Order Rhizomastigida
Class Dictyostelea
Order Dictyostelida
Class Myxogastrea [= Myxomycetes]
Subclass Exosporeae
Order Ceratiomyxida
Subclass Myxogastria
Superorder Columelida
Order Echinosteliida
Order Fuscisporida
Superorder Lucisporida
Order Liceida
Order Trichiida
Class Protostelea
Order Protostelida
Class Variosea
Order Artodiscida
Order Holomastigida
Order Phalansteriida
Order Varipodida
Subphylum Lobosa
Class Discosea
Subclass Flabellinia
Order Dactylopodida
Order Himatismenida
Order Pellitida
(Continued)
16 / 60
Order Stygamoebida
Order Trichosida
Order Vanellida
Subclass Longamoebia
Order Dermamoebida
Order Centramoebida
Order Thecamoebida
Class Tubulinea [= Lobosea]
Order Arcellinida
Order Echinamoebida
Order Euamoebida
Order Leptomyxida
Order Nolandida
Phylum Choanozoa [with Microsporidia, Animalia, and Fungi constitutes "Supergroup Opisthokonta"]
Subphylum Choanola
Class Choanoagellatea
Order Acanthoecida
Order Craspedida
Class Corallochytrea
Order Corallochytrida
Class Filasterea
Order Ministeriida
Class Ichthyosporea
Order Dermocystida
Order Eccrinida
Subphylum Paramycia
Class Aphelidea
Order Aphelidida
Class Cristidiscoidea
Order Fonticulida
Order Nucleariida
Class Rozellidea
Order Rozellida
Phylum Microsporidia [with Choanozoa, Animalia, and Fungi constitutes "Supergroup Opisthokonta"]
Class Disporea
Order N.N. (e.g., Nosema)
Class Metchnikovellea
Order Metchnikovellida
Class Minisporea [= Microsporea]
Order Minisporida [= Minisporea]
Class Pleistophorea
Order Pleistophorida
Phylum Sulcozoa
Subphylum Apusozoa
Class Breviatea
Order Breviatida
Class Thecomonadea
Order Apusomonadida
Subphylum Varisulca
Class Diphyllatea
(Continued)
17 / 60
Order Diphylleida
Class Glissodiscea
Order Mantamonadida
Order Planomonadida
Class Hilomonadea
Order Rigilida
KINGDOM CHROMISTA
SUBKINGDOM HACROBIA
Phylum N.N.
Class Endohelea
Order Heliomonadida
Order Microhelida
Class Picomonadea
Order Picomonadida
Class Telonemea
Order Telonemida
Phylum Cryptista
Subphylum Palpitia
Class Palpitea
Subphylum Rollomonadia
Class Cryptophyceae
Order Cryptomonadales
Order Pyrenomonadales
Order Tetragonidiales
Class Goniomonadea
Order Goniomonadida
Class Leucocryptea
Order Katablepharida
Order Palpitida
Phylum Haptophyta
Class Coccolithophyceae [= Prymnesiophyceae]
Order Coccolithales
Order Coccosphaerales
Order Isochrysidales
Order Phaeocystales
Order Prymnesiales
Order Syracosphaerales
Order Zygodiscales
Class Pavlovophyceae
Order Pavlovales
Phylum Heliozoa
Class Centrohelea
Order Acanthocystida
Order Pterocystida
SUBKINGDOM HAROSA [= "Supergroup SAR"]
INFRAKINGDOM HALVARIA
Superphylum Alveolata
Phylum Ciliophora
Subphylum Intramacronucleata
Class Armophorea
(Continued)
18 / 60
Order Armophorida
Order Clevelandellida
Class Colpodea
Order Bryometopida
Order Bryophryida
Order Bursariomorphida
Order Colpodida
Order Cyrtolophosidida
Order Sorogenida
Class Litostomatea
Subclass Haptoria
Order Cyclotrichiida
Order Haptorida
Order Pleurostomatida
Subclass Trichostomatia
Order Entodiniomorphida
Order Macropodiniida
Order Vestibuliferida
Class Spirotrichea
Subclass Choreotrichia
Order Tintinnida
Subclass Hypotrichia
Order Euplotida
Order Kiitrichida
Subclass Licnophoria
Order Licnophorida
Subclass Oligotrichia
Order Strombidiida
Subclass Protocruziidia
Order Phacodiniida
Order Protocruziida
Subclass Stichotrichia
Order Sporadotrichida
Order Stichotrichida
Order Urostylida
Class Nassophorea
Order Colpodidiida
Order Microthoracida
Order Nassulida
Order Synhymeniida
Class Oligohymenophorea
Subclass Apostomatia
Order Apostomatida
Order Astomatophorida
Order Pilisuctorida
Subclass Astomatia
Order Astomatida
Subclass Hymenostomatia
Order Ophyroglenida
Order Tetrahymenida
(Continued)
19 / 60
Subclass Peniculia
Order Peniculida
Order Urocentrida
Subclass Peritrichia
Order Mobilida
Order Sessilida
Subclass Scuticociliatia
Order Philasterida
Order Pleuronematida
Order Thigmotrichida
Class Phyllopharyngea
Subclass Chonotrichia
Order Cryptogemmida
Order Exogemmiida
Subclass Cyrtophoria
Order Chlamydodontida
Order Dysteriida
Subclass Rhynchodia
Order Hypocomatida
Order Rhynchodida
Subclass Suctoria
Order Endogenida
Order Evaginogenida
Order Exogenida
Class Plagiopylea
Order Odontostomatida
Order Plagiopylida
Class Prostomatea
Order Prorodontida
Order Prostomatida
Subphylum Postciliodesmatophora
Class Heterotrichea
Order Heterotrichida
Class Karyorelictea
Order Loxodida
Order Protoheterotrichida
Order Protostomatida
Phylum Miozoa
Subphylum Myzozoa
Infraphylum Apicomplexa
Superclass Apicomonada
Class Apicomonadea
Order Chromerida
Order Colpodellida
Order Voromonadida
Superclass Sporozoa
Class N.N.
Order Blastogregarinida
Class Coccidiomorphea
Subclass Coccidea
(Continued)
20 / 60
Order Agamococcidida
Order Eimerida
Order Ixorheida
Subclass Coleotrophia
Order Coleotrophiida [= Protococciida]
Subclass Hematozoa
Superorder Aconoidia
Order Nephromycida
Order Piroplasmida
Superorder Haemosporidia
Order Hemosporida
Class Gregarinomorphea
Subclass Cryptogragaria
Order Cryptogregarida (Cryptosporidium)
Subclass Histogregaria
Order Histogregarida
Subclass Orthogregarinia
Order Arthrogarida
Order Vermigregarida
Class Paragregarea
Order Archigregarinida
Order Stenophorida
Order Velocida
Infraphylum Dinozoa
Superclass Dinoagellata
Class Dinophyceae
Subclass N.N.
Order Actiniscales
Order Blastodinales
Order Coccidinales
Order Dinamoebales
Order Lophodinales
Order Pyrocystales
Order Thoracosphaerales
Subclass Dinophysoidia
Order Dinophysidales
Order Nannoceratopsales
Subclass Gonyaulacoidia
Order Gonyaulacales
Order Gymnodiniales
Subclass Peridinoidia
Order Peridiniales
Order Prorocentrales
Subclass Suessioidia
Order Suessiales
Class Ellobiopsea
Order Ellobiopsida
Class Noctilucea
Order Noctilucida
Class Oxyrrhea
(Continued)
21 / 60
Order Acrocoelida
Order Oxyrrhida
Class Syndinea
Order Rastrimonadida
Order Syndinida
Superclass Perkinsozoa
Class Myzomonadea
Order Algovorida
Class Perkinsea
Order Perkinsida
Order Phagodinida
Subphylum Protalveolata
Class Colponemea
Order Colponemida
Superphylum Heterokonta [= "Supergroup Stramenopiles"]
Phylum Bigyra
Class Bikosea
Order Anoecida
Order Bicoecida
Order Borokida
Order Pseudodendromonadida
Order Rictida
Class Blastocystea
Order Blastocystida
Class Nanomonadea
Order Uniciliatida
Class Opalinea
Order Opalinida
Order Proteromonadida
Class Labyrinthulea
Order Labyrinthulida
Order Thraustochytriida
Class Placididea [= Placidiophyceae]
Order Placidiida
Phylum Ochrophyta [= Heterokontophyta p.p.]
Class Bacillariophyceae [= Diatomeae]
Subclass Bacillariophycidae
Order Achnanthales
Order Bacillariales
Order Cymbellales
Order Dictyoneidales
Order Eunotiales
Order Eupodiscales
Order Lyrellales
Order Mastogloiales
Order Naviculales
Order Rhopalodiales
Order Surirellales
Order Thalassiophysales
Subclass Coscinodiscophycidae
(Continued)
22 / 60
Order Anaulales
Order Arachnoidiscales
Order Asterolamprales
Order Aulacoseirales
Order Biddulphiales
Order Chaetocerotales
Order Chrysanthemodiscales
Order Corethrales
Order Coscinodiscales
Order Cymatosirales
Order Ethmodiscales
Order Hemiaulales
Order Lithodesmiales
Order Melosirales
Order Orthoseirales
Order Paraliales
Order Rhizosoleniales
Order Stictocyclales
Order Stictodiscales
Order Thalassiosirales
Order Triceratiales
Subclass Fragilariophycidae
Order Ardissoneales
Order Climacospheniales
Order Cyclophorales
Order Fragilariales
Order Licmophorales
Order Protoraphidales
Order Rhabdonematales
Order Rhaphoneidales
Order Striatellales
Order Tabellariales
Order Thalassionematales
Order Toxariales
Class Bolidophyceae
Order Parmales [= Bolidomonadales]
Class Chrysomerophyceae
Order Chrysomeridales
Class Chrysophyceae
Order Chloramoebales
Order Chromulinales
Order Chrysosphaerales
Order Heterogloeales
Order Hibberdiales
Order Hydrurales
Order Ochromonadales
Order Paraphysomonadida
Order Synurales
Order Thallochrysidales
Class Eustigmatophyceae
(Continued)
23 / 60
Order Eustigmatales
Class Dictyochophyceae [= Hypogyristea]
Order Dictyochales
Order Olisthodiscales
Order Pedinellales
Order Pelagomonadales
Order Sarcinochrysidales
Class Phaeophyceae
Subclass Dictypophycidae
Order Dictyotales
Order Onslowiales
Order Sphacelariales
Order Syringodermatales
Subclass Discosporangiophycidae
Order Discosporangiales
Subclass Fucophycidae
Order Ascoseirales
Order Asterocladales
Order Desmarestiales
Order Ectocarpales
Order Fucales
Order Laminariales
Order Nemodermatales
Order Phaeosiphoniellales
Order Ralfsiales
Order Scytothamnales
Order Sporochnales
Order Tilopteridales [= Cutleriales]
Subclass Ishigeophycidae
Order Ishigeales
Class Phaeothamniophyceae [= Aurophyceae]
Order Aurearenales
Order Phaeothamniales
Class Picophagophyceae [= Picophagea]
Order Picophagales
Order Synchromales
Class Pinguiophyceae
Order Pinguiochrysidales
Class Raphidophyceae
Order Actinophryida
Order Commatiida
Order Raphidomonadales
Class Schizocladiophyceae
Order Schizocladiales
Class Xanthophyceae
Order Mischococcales
Order Pleurochloridellales
Order Tribonematales
Order Vaucheriales
Phylum Pseudofungi [= Oomycota]
(Continued)
24 / 60
Class Bigyromonadea
Order Developayellida
Class Hyphochytrea
Order Hyphochytriida
Order Pirsoniida
Class Oomycetes
Subclass Eogamia
Order Anisolpidiales
Order Haptoglossales
Order Lagenismatales
Order Olpidiopsidales
Order Rozellopsidales
Subclass Peronosporidae
Order Peronosporales
Order Pythiales
Order Rhipidiales
Subclass Saprolegniidae
Order Albuginales
Order Leptomitales
Order Salilagenidiales
Order Saprolegniales
INFRAKINGDOM RHIZARIA
Phylum Cercozoa
Subphylum Endomyxa
Class Ascetosporea
Order Claustrosporida
Order Haplosporida
Order Paradinida
Order Paramyxida
Class Gromiidea
Order Gromiida
Order Reticulosida
Class Phytomyxea
Order Phagomyxida
Order Plasmodiophorida
Class Vampyrellidea
Order Vampyrellida
Subphylum Monadolosa
Class Imbricatea
Subclass Placonuda
Order Discocelida
Order Discomonadida
Order Euglyphida
Order Marimonadida
Order Variglissida
Subclass Placoperla
Order Perlolida
Order Rotosphaerida
Order Spongomonadida
Order Thaumatomonadida
(Continued)
25 / 60
Order Zoelucasida
Class Metromonadea
Order Metopiida
Order Metromonadida
Class Sarcomonadea
Order Cercomonadida
Order Glissomonadida
Order Pansomonadida
Order Pseudosporida
Order Sainouroida
Class Thecolosea
Subclass Eothecia
Order Cryomonadida
Order Ebriida
Order Matazida
Order Ventricleftida
Subclass Phaeodaria
Order Eodarida
Order Opaloconchida
Subclass Tectosia
Order Tectolosida
Subphylum Reticulolosa
Class Chlorarachnea
Order Chlorarachnida
Class Granolosea
Order Cryptolida
Order Desmothoracida
Order Leucodictyida
Order Limnolida
Class Skiomonadea
Order Tremulida
Phylum Retaria
Subphylum Foraminifera
Class Monothalamea
Order Allogromiida
Order Astrorhizida
Order Psamminida
Order Stannomida
Class Globothalamea
Order Carterinida
Order Globeriginida
Order Lagenida
Order Lituolida
Order Lofusiida
Order Robertinida
Order Rotaliida
Order Testulariida
Order Trochamminida
Class Tubothalamea
Order Miliolida
(Continued)
26 / 60
Order Spirillinida
Subphylum Radiozoa
Superclass Polycystinia
Class Polycystinea
Order Collodarida
Order Nassellaria
Order Spumellaria
Superclass Spasmaria
Class Acantharea
Order Arthracanthida
Order Chaunacanthida
Order Holacanthida
Order Symphyacanthida
Class Sticholonchea
Order Taxopodida
KINGDOM FUNGI
SUBKINGDOM DIKARYA [= NEOMYCOTA]
Phylum Ascomycota
Subphylum Pezizomycotina
Class Archaeorhizomycetes
Order Archaeorhizomycetales
Order Lahmiales
Order Triblidiales
Class Arthoniomycetes
Order Arthoniales
Class Dothideomycetes
Subclass N.N.
Order Acrospermales
Order Botryosphaeriales
Order Hysteriales
Order Jahnulales
Order Koralionastetales
Order Patellariales
Order Trypetheliales
Subclass Dothideomycetidae
Order Capnodiales
Order Dothideales
Order Microthyriales
Order Myriangiales
Subclass Meliolomycetidae
Order Meliolales
Subclass Pleosporomycetidae
Order Mytilinidales
Order Pleosporales
Class Eurotiomycetes
Subclass Chaetothyriomycetidae
Order Chaetothyriales
Order Pyrenulales
Order Verrucariales
Subclass Eurotiomycetidae
(Continued)
27 / 60
Order Arachnomycetales
Order Ascosphaerales
Order Coryneliales
Order Eurotiales
Order Onygenales
Subclass Mycocaliciomycetidae
Order Mycocaliciales
Class Laboulbeniomycetes
Order Laboulbeniales
Order Pyxidiophorales
Class Lecanoromycetes
Subclass N.N.
Order Candelariales
Order Umbilicariales
Subclass Acarosporomycetidae
Order Acarosporales
Subclass Lecanoromycetidae
Order Lecanorales
Order Lecideales
Order Peltigerales
Order Rhizocarpales
Order Teloschistales
Subclass Ostropomycetidae
Order Agyriales
Order Baeomycetales
Order Ostropales
Order Pertusariales
Class Leotiomycetes
Order Cyttariales
Order Erysiphales
Order Geoglossales
Order Helotiales
Order Leotiales
Order Mediolariales
Order Rhytismatales
Order Thelebolales
Class Lichinomycetes
Order Eremithallales
Order Lichinales
Class Orbiliomycetes
Order Orbiliales
Class Pezizomycetes
Order Pezizales
Class Sordariomycetes
Subclass N.N.
Order Phyllachorales
Order Trichosphaeriales
Subclass Hypocreomycetidae
Order Coronophorales
Order Hypocreales
(Continued)
28 / 60
Order Melanosporales
Order Microascales
Subclass Sordariomycetidae
Order Boliniales
Order Calosphaeriales
Order Chaetosphaeriales
Order Coniochaetales
Order Diaporthales
Order Ophiostomatales
Order Sordariales
Subclass Spathulosporomycetidae
Order Lulworthiales
Subclass Xylariomycetidae
Order Xylariales
Subphylum Saccharomycotina
Class Saccharomycetes
Order Saccharomycetales
Subphylum Taphrinomycotina
Class Neolectomycetes
Order Neolectales
Class Pneumocystidomycetes
Order Pneumocystidales
Class Schizosaccharomycetes
Order Schizosaccharomycetales
Class Taphrinomycetes
Order Taphrinales
Phylum Basidiomycota
Class Entorrhizomycetes
Order Entorrhizales
Order Wallemiales
Subphylum Agaricomycotina
Class Agaricomycetes
Subclass N.N.
Order Auriculariales
Order Cantharellales
Order Corticiales
Order Gloeophyllales
Order Hymenochaetales
Order Polyporales
Order Russulales
Order Sebacinales
Order Thelephorales
Order Trechisporales
Subclass Agaricomycetidae
Order Agaricales
Order Atheliales
Order Boletales
Subclass Phallomycetidae
Order Geastrales
Order Gomphales
(Continued)
29 / 60
Order Hysterangiales
Order Phallales
Class Dacrymycetes
Order Dacrymycetales
Class Tremellomycetes
Order Cystolobasidiales
Order Filobasidiales
Order Tremellales
Subphylum Pucciniomycotina
Class Agaricostilbomycetes
Order Agaricostilbales
Order Spiculogloeales
Class Atractiellomycetes
Order Atractiellales
Class Classiculomycetes
Order Classiculales
Class Cryptomycocolacomycetes
Order Cryptomycocolacales
Class Cystobasidiomycetes
Order Cystobasidiales
Order Erythrobasidiales
Order Naohideales
Class Microbotryomycetes
Order Hetrogastridiales
Order Leucosporidiales
Order Microbotryales
Order Sporidiobolales
Class Mixiomycetes
Order Mixiales
Class Pucciniomycetes
Order Helicobasidiales
Order Pachnocybales
Order Platygloeales
Order Pucciniales
Order Septobasidiales
Subphylum Ustilaginomycotina
Class N.N.
Order Malasseziales
Class Exobasidiomycetes
Order Ceraceosorales
Order Doassansiales
Order Entylomatales
Order Exobasidiales
Order Georgescheriales
Order Microstromatales
Order Tilletiales
Class Ustilaginomycetes
Order Urocystidales
Order Ustilaginales
SUBKINGDOM EOMYCOTA
(Continued)
30 / 60
Phylum Chytridiomycota
Class Blastocladiomycetes [= Allomycetes]
Order Blastocladiales
Class Chytridiomycetes
Order Chytridiales
Order Lobulomycetales
Order Neocallimastigales
Order Olpidiales
Order Rhizophlyctidales
Order Rhizophydiales
Order Spizellomycetales
Class Monoblepharidomycetes
Order Monoblepharidales
Phylum Glomeromycota
Class Glomeromycetes [= Glomomycetes]
Order Archaeosporales
Order Diversisporales
Order Glomerales
Order Paraglomerales
Phylum Zygomycota
Subphylum N.N.
Class N.N.
Order Basidiobolales
Subphylum Entomophthoromycotina
Class N.N.
Order Entomophthorales
Subphylum Kickxellomycotina
Class N.N.
Order Asellariales
Order Dimargaritales
Order Harpellales
Order Kickxellales
Subphylum Mortierellomycotina
Class N.N.
Order Mortierellales
Subphylum Mucoromycotina
Class N.N.
Order Endogonales
Order Mucorales
Subphylum Zoopagomycotina
Class N.N.
Order Zoopagales
KINGDOM PLANTAE
SUBKINGDOM BILIPHYTA
Phylum Glaucophyta
Class Glaucophyceae
Order Glaucocystales
Phylum Rhodophyta
Subphylum Cyanidiophytina
Class Cyanidiophyceae
(Continued)
31 / 60
Order Cyanidiales
Subphylum Eurhodophytina
Class Bangiophyceae
Order Bangiales
Order Goniotrichales
Class Florideophyceae
Subclass N.N.
Order Rhodachlyales
Subclass Ahnfeltiophycidae
Order Ahnfeltiales
Order Pihiellales
Subclass Corallinophycidae
Order Corallinales
Order Rhodogorgonales
Order Sporolithales
Subclass Hildenbrandiophyceae
Order Hildenbrandiales
Subclass Nemaliophycidae
Order Acrochaetiales
Order Balbianiales
Order Balliales
Order Batrachospermales
Order Colaconematales
Order Entwisleiales
Order Nemaliales
Order Palmariales
Order Thoreales
Subclass Rhodymeniophycidae
Order Acrosymphytales
Order Bonnemaisoniales
Order Ceramiales
Order Gelidiales
Order Gigartinales
Order Gracilariales
Order Halymeniales
Order Nemastomatales
Order Peyssonneliales
Order Plocamiales
Order Rhodymeniales
Order Sebdeniales
Subphylum Metarhodophytina
Class Compsopogonophyceae
Order Compsopogonales
Order Erythropeltidales
Order Rhodochaetales
Subphylum Rhodellophytina
Class Porphyridiophyceae
Order Porphyridiales
Class Rhodellophyceae
Order Dixoniellales
(Continued)
32 / 60
Order Glaucosphaerales
Order Rhodellales
Class Stylonematophyceae
Order Rufusiales
Order Stylonematales
SUBKINGDOM VIRIDIPLANTAE
INFRAKINGDOM CHLOROPHYTA
Phylum Chlorophyta
Subphylum Chlorophytina
Class Chlorodendrophyceae
Order Chlorodendrales
Class Chlorophyceae
Order N.N. (e.g., Chlorangiopsidaceae)
Order Chaetopeltidales
Order Chaetophorales
Order Chlamydomonadales [= Volvocales]
Order Oedogoniales
Order Sphaeropleales
Class Pedinophyceae
Order Marsupiomonadales
Order Pedinomonadales
Order Scoureldiales
Class Trebouxiophyceae
Order Chlorellales
Order Microthamniales
Order Phyllsiphonales
Order Prasiolales
Order Trebouxiales
Class Ulvophyceae
Order Bryopsidales
Order Cladophorales
Order Dasycladales
Order Oltmansiellopsidales
Order Scotinosphaerales
Order Trentepohliales
Order Ulotrichales
Order Ulvales
Subphylum Prasinophytina
Class Mamiellophyceae
Order Dolichomastigales
Order Mamiellales
Order Monomastigales
Class Nephrophyceae [= Nephroselmidophyceae]
Order Nephroselmidales
Class Pyramimonadophyceae
Order Palmophyllales
Order Prasinococcales
Order Pseudoscoureldiales
Order Pyramimonadales
INFRAKINGDOM STREPTOPHYTA
(Continued)
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Superphylum Charophyta
Phylum Charophyta
Class Charophyceae
Order Charales
Class Chlorokybophyceae
Order Chlorokybales
Class Coleochaetophyceae
Order Chaetosphaeridiales
Order Coleochaetales
Class Conjugatophyceae [= Zygnematophyceae]
Order Desmidiales
Order Zygnematales
Class Klebsormidiophyceae
Order Klebsormidiales
Class Mesostigmatophyceae
Order Mesostigmatales
Superphylum Embryophyta
Phylum Anthocerotophyta
Class Anthocerotopsida
Subclass Anthocerotidae
Order Anthocerotales
Subclass Dendrocerotidae
Order Dendrocerotales
Order Phymatocerales
Subclass Notothylatidae
Order Notothyladales
Class Leiosporocerotopsida
Order Leiosporocerotales
Phylum Bryophyta
Class Andreaeobryopsida
Order Andreaeobryales
Class Andreaeopsida
Order Andreaeales
Class Bryopsida
Subclass Bryidae
Order Bartramiales
Order Bryales
Order Hedwigiales
Order Hookeriales
Order Hypnales
Order Hypnodendrales
Order Orthotrichales
Order Ptychomniales
Order Rhizogoniales
Order Splachnales
Subclass Buxbaumiidae
Order Buxbaumiales
Subclass Dicranidae
Order Archidiales
Order Bryoxiphiales
(Continued)
34 / 60
Order Dicranales
Order Grimmiales
Order Pottiales
Order Scouleriales
Subclass Diphysciidae
Order Diphysciales
Subclass Funariidae
Order Encalyptales
Order Funariales
Order Gigaspermales
Subclass Timmidae
Order Timmiales
Class Oedipodiopsida
Order Oedipodiales
Class Polytrichopsida
Order Polytrichales
Class Sphagnopsida
Order Ambuchananiales
Order Sphagnales
Class Takakiopsida
Order Takakiales
Class Tetraphidopsida
Order Tetraphidales
Phylum Marchantiophyta
Class Haplomitriopsida
Order Calobryales
Order Treubiales
Class Jungermanniopsida
Subclass Jungermanniidae
Order Jungermanniales
Order Porellales
Order Ptilidiales
Subclass Metzgeriidae
Order Metzgeriales
Order Pleuroziales
Subclass Pelliidae
Order Fossombroniales
Order Pallaviciniales
Order Pelliales
Class Marchantiopsida
Order Blasiales
Order Lunulariales
Order Marchantiales
Order Neohodgsoniales
Order Sphaerocarpales
Phylum Tracheophyta
Subphylum Lycopodiophytina
Class Lycopodiopsida
Order Isoetales
Order Lycopodiales
(Continued)
35 / 60
Order Selaginellales
Subphylum Polypodiophytina
Class Polypodiopsida
Subclass Equisetidae
Order Equisetales
Subclass Marattiidae
Order Marattiales
Subclass Ophioglossidae [= Psilotidae]
Order Ophioglossales
Order Psilotales
Subclass Polypodiidae
Order Cyatheales
Order Gleicheniales
Order Hymenophyllales
Order Osmundales
Order Polypodiales
Order Salviniales
Order Schizaeales
Subphylum Spermatophytina
Superclass Angiospermae
Class Magnoliopsida
Superorder N.N.
Order N.N. (e.g., Icacinaceae)
Superorder Amborellanae
Order Amborellales
Superorder Asteranae
Order Apiales
Order Aquifoliales
Order Asterales
Order Boraginales
Order Bruniales
Order Cornales
Order Dipsacales
Order Ericales
Order Escalloniales
Order Garryales
Order Gentianales
Order Lamiales
Order Paracryphiales
Order Solanales
Superorder Austrobaileyanae
Order Austrobaileyales
Superorder Berberidopsidanae
Order Berberidopsidales
Superorder Buxanae
Order Buxales
Superorder Caryophyllanae
Order Caryophyllales
Superorder Ceratophyllanae
Order Ceratophyllales
(Continued)
36 / 60
Superorder Dillenianae
Order Dilleniales
Superorder Lilianae [= Monocotyledones]
Order Acorales
Order Alismatales
Order Arecales
Order Asparagales
Order Commelinales
Order Dasypogonales
Order Dioscoreales
Order Liliales
Order Pandanales
Order Petrosaviales
Order Poales
Order Zingiberales
Superorder Magnolianae
Order Canellales
Order Chloranthales
Order Laurales
Order Magnoliales
Order Piperales
Superorder Myrothamnanae
Order Gunnerales
Superorder Nymphaeanae
Order Nymphaeales
Superorder Proteanae
Order Proteales
Superorder Ranunculanae
Order Ranunculales
Superorder Rosanae
Order Brassicales
Order Celastrales
Order Crossosomatales
Order Cucurbitales
Order Fabales
Order Fagales
Order Geraniales
Order Huerteales
Order Malpighiales
Order Malvales
Order Myrtales
Order Oxalidales
Order Picramniales
Order Rosales
Order Sapindales
Order Vitales
Order Zygophyllales
Superorder Santalanae
Order Santalales
Superorder Saxifraganae
(Continued)
37 / 60
Order Saxifragales
Superorder Trochodendranae
Order Trochodendrales
Superclass Gymnospermae
Class Cycadopsida
Subclass Cycadidae
Order Cycadales
Class Ginkgoopsida
Subclass Ginkgooidae
Order Ginkgoales
Class Gnetopsida
Subclass Gnetidae
Order Gnetales
Class Pinopsida
Subclass Pinidae
Order Pinales
KINGDOM ANIMALIA
SUBKINGDOM N.N.
Phylum Cnidaria
Subphylum Anthozoa
Class Anthozoa
Subclass Hexacorallia
Order Actiniaria
Order Antipatharia
Order Ceriantharia
Order Corallimorpharia
Order Scleractinia
Order Zoantharia [= Zoanthidea]
Subclass Octocorallia
Order Alcyonacea
Order Helioporacea
Order Pennatulacea
Subphylum Medusozoa
Class Cubozoa
Order Carybdeida
Order Chirodropida
Class Hydrozoa
Subclass Hydroidolina
Order Anthoathecata
Order Gonoproxima
Order Leptothecata
Order Siphonophorae
Subclass Trachylina
Order Actinulida
Order Limnomedusae
Order Narcomedusae
Order Trachymedusae
Class Polypodiozoa
Order Polypodiidea
Class Scyphozoa
(Continued)
38 / 60
Order Coronatae
Order Rhizostomeae
Order Semaeostomeae
Class Staurozoa
Order Stauromedusae
Subphylum Myxozoa
Class Malacosporea
Order Malacovalvulida
Class Myxosporea
Order Bivalvulida
Order Multivalvulida
Phylum Ctenophora
Class Nuda
Order Beroida
Class Tentaculata
Order Cambojiida
Order Cestida
Order Cryptolobiferida
Order Cydippida
Order Ganeshida
Order Lobata
Order Platyctenida
Order Thalassocalycida
Phylum Placozoa
Class Placozoa (Trichoplax)
Phylum Porifera
Class Calcarea
Order Baerida
Order Clathrinida
Order Leucosolenida
Order Lithonida
Order Murrayonida
Class Demospongiae
Order Agelasida
Order Astrophorida
Order Chondrosida
Order Dendroceratida
Order Dictyoceratida
Order Hadromerida
Order Halichondrida
Order Haplosclerida
Order Lithistida
Order Poecilosclerida
Order Spirophorida
Order Verongida
Class Hexactinellida
Order Amphidiscosida
Order Aulocalycoida
Order Fieldingida
Order Hexactinosida
(Continued)
39 / 60
Order Lychniscosida
Order Lyssacinosida
Class Homoscleromorpha
Order Homosclerophorida
SUBKINGDOM BILATERIA
INFRAKINGDOM PROTOSTOMIA
Superphylum N.N.
Phylum Chaetognatha
Class Sagittoidea
Order Aphragmophora
Order Phragmophora
Phylum Orthonectida
Order Plasmodigenea
Phylum Rhombozoa
Order Dicyemida
Order Heterocyemida
Superphylum Ecdysozoa
Phylum Arthropoda
Subphylum Chelicerata
Class Arachnida
Superorder N.N.
Order Amblypygi
Order Araneae
Order Opiliones
Order Palpigradi
Order Pseudoscorpiones
Order Ricinulei
Order Schizomida
Order Scorpiones
Order Solifugae
Order Uropygi
Superorder Acariformes
Order Sarcoptiformes
Order Trombidiformes
Superorder Parasitiformes
Order Holothyrida
Order Ixodida
Order Mesostigmata
Order Opilioacarida
Class Merostomata
Order Xiphosura
Class Pycnogonida
Order Pantopoda
Subphylum Crustacea
Class Branchiopoda
Order Anostraca
Order Diplostraca
Order Laevicaudata
Order Notostraca
Class Cephalocarida
(Continued)
40 / 60
Order Brachypoda
Class Malacostraca
Subclass Eumalacostraca
Superorder Eucarida
Order Amphionidacea
Order Decapoda
Order Euphausiacea
Superorder Peracarida
Order Amphipoda
Order Bochusacea
Order Cumacea
Order Isopoda
Order Lophogastrida
Order Mictacea
Order Mysida
Order Spelaeogriphacea
Order Tanaidacea
Order Thermosbaenacea
Superorder Syncarida
Order Anaspidacea
Order Bathynellacea
Subclass Hoplocarida
Order Stomatopoda
Subclass Phyllocarida
Order Leptostraca
Class Maxillopoda
Subclass Branchiura
Order Arguloida
Subclass Copepoda
Infraclass Neocopepoda
Superorder Gymnoplea
Order Calanoida
Superorder Podoplea
Order Cyclopoida
Order Gelyelloida
Order Harpacticoida
Order Misophrioida
Order Monstrilloida
Order Mormonilloida
Order Siphonostomatoida
Infraclass Progymnoplea
Order Platycopioida
Subclass Mystacocarida
Order Mystacocaridida
Subclass Pentastomida
Order Cephalobaenida
Order Porocephalida
Subclass Tantulocarida (e.g., Basipodellidae)
Subclass Thecostraca
Infraclass Ascothoracida
(Continued)
41 / 60
Order Dendrogastrida
Order Laurida
Infraclass Cirripedia
Superorder Acrothoracica
Order Cryptophialida
Order Lithoglyptida
Superorder Rhizocephala
Order Akentrogonida
Order Kentrogonida
Superorder Thoracica
Order Ibliformes
Order Lepadiformes
Order Scalpelliformes
Order Sessilia
Infraclass Facetotecta (Hansenocaris)
Class Ostracoda
Order Halocyprida
Order Myodocopida
Order Paleocopida
Order Platycopida
Order Podocopida
Class Remipedia
Order Nectiopoda
Subphylum Hexapoda
Class Collembola
Order Entomobryomorpha
Order Neelipleona
Order Poduromorpha
Order Symphypleona
Class Diplura
Order N.N. (e.g., Japygidae)
Class Insecta
Subclass Archaeognatha
Order Archaeognatha
Subclass Dicondylia
Order Zygentoma
Subclass Pterygota
Infraclass Neoptera
Superorder Holometabola
Order Coleoptera
Order Diptera
Order Hymenoptera
Order Lepidoptera
Order Mecoptera
Order Siphonaptera
Order Strepsiptera
Order Trichoptera
Superorder Neuropterida
Order Megaloptera
Order Neuroptera
(Continued)
42 / 60
Order Raphidioptera
Superorder Paraneoptera
Order Hemiptera
Order Psocodea
Order Thysanoptera
Superorder Polyneoptera
Order Blattodea
Order Dermaptera
Order Embioptera
Order Grylloblattodea
Order Mantodea
Order Mantophasmatodea
Order Orthoptera
Order Phasmida
Order Plecoptera
Order Zoraptera
Infraclass Palaeoptera
Order Ephemeroptera
Order Odonata
Class Protura
Order Acerentomata
Order Eosentomata
Order Sinentomata
Subphylum Myriapoda
Class Chilopoda
Order Craterostigmomorpha
Order Geophilomorpha
Order Lithobiomorpha
Order Scolopendromorpha
Order Scutigeromorpha
Class Diplopoda
Subclass Chilognatha
Infraclass Helminthomorpha
Superorder N.N.
Order Platydesmida
Order Polyzoniida
Order Siphonocryptida
Order Siphonophorida
Superorder Juliformia
Order Julida
Order Spirobolida
Order Spirostreptida
Superorder Nematophora
Order Callipodida
Order Chordeumatida
Order Stemmiulida
Order Siphoniulida
Superorder Merochaeta
Order Polydesmida
Infraclass Pentazonia
(Continued)
43 / 60
Order Glomerida
Order Glomeridesmida
Order Sphaerotheriida
Subclass Penicillata
Order Polyxenida
Class Pauropoda
Order Hexamerocerata
Order Tetramerocerata
Class Symphyla (e.g., Scolopendrellidae)
Phylum Kinorhyncha
Order Cyclorhagida
Order Homalorhagida
Phylum Loricifera
Order Nanaloricida
Phylum Nematoda
Class Chromadorea
Subclass Chromadoria
Order Chromadorida
Order Desmodorida
Order Desmoscolecida
Order Selachinematida
Subclass Plectia
Superorder Monhysterica
Order Monhysterida
Superorder Plectica
Order Benthimermithida
Order Leptolaimida
Order Plectida
Superorder Rhabditica
Order Diplogasterida
Order Drilonematida
Order Panagrolaimida
Order Rhabditida
Order Spirurida
Superorder Teratocephalica
Order Teratocephalida
Class Dorylaimea
Subclass Bathyodontia
Order Bathyodontida
Order Mermithida
Order Mononchida
Subclass Dorylaimia
Order Dorylaimida
Subclass Trichocephalia
Order Dioctophymatida
Order Marimermithida
Order Muspiceida
Order Trichocephalida
Class Enoplea
Subclass Enoplia
(Continued)
44 / 60
Order Alaimida
Order Enoplida
Order Ironida
Order Rhaptothyreida
Order Trifusiida
Order Tripyloidida
Subclass Oncholaimia
Order Oncholaimida
Subclass Triplonchia
Order Triplonchida
Order Tripylida
Phylum Nematomorpha
Order Gordioidea
Order Nectonematoidea
Phylum Onychophora
Class Udeonycophora
Order Euonycophora
Phylum Priapula [= Priapulida]
Class N.N. (e.g., Priapulidae)
Phylum Tardigrada
Class Eutardigrada
Order Apochela
Order Parachela
Class Heterotardigrada
Order Arthrotardigrada
Order Echiniscoidea
Superphylum Spiralia [= Lophotrochozoa]
Phylum Acanthocephala
Class Archiacanthocephala
Order Apororhynchida
Order Gigantorhynchida
Order Moniliformida
Order Oligacanthorhynchida
Class Eoacanthocephala
Order Gyracanthocephala
Order Neoechinorhynchida
Class Palaeacanthocephala
Order Echinorhynchida
Order Heteramorphida
Order Polymorphida
Class Polyacanthocephala
Order Polyacanthorhynchida
Phylum Annelida
Class N.N.
Order Myzostomida
Class Clitellata
Subclass N.N.
Order Apodadrilida
Subclass Hirudinea
Order Acanthobdellida
(Continued)
45 / 60
Order Arhynchobdellida
Order Rhynchobdellida
Subclass Oligochaeta
Superorder N.N.
Order N.N. (Jennaria)
Order Branchiobdellida
Order Capilloventrida
Order Crassiclitellata
Order Enchytraeida
Order Haplotaxida
Order Lumbriculida
Order Tubicida
Superorder Metagynaphora
Order Moniligastrida
Order Opistophophora
Class Polychaeta
Subclass N.N.
Order N.N. (e.g., Nerillidae)
Subclass Echiura
Order Echiuroinea
Order Heteromyota
Order Xenopneusta
Subclass Errantia
Order Amphinomida
Order Eunicida
Order Phyllodocida
Subclass Sedentaria
Infraclass Canalipalpata
Order Sabellida
Order Spionida
Order Terebellida
Infraclass Scolecida (e.g., Arenicolidae)
Phylum Brachiopoda
Class Craniata
Order Craniida
Class Lingulata
Order Lingulida
Class Rhynchonellata
Order Rhynchonellida
Order Terebratulida
Order Thecideida
Phylum Bryozoa
Class Gymnolaemata
Order Cheilostomata
Order Ctenostomata
Class Phylactolaemata
Order Plumatellida
Class Stenolaemata
Order Cyclostomata
Phylum Cycliophora
(Continued)
46 / 60
Class Eucycliophora
Order Symbiida
Phylum Entoprocta
Order Coloniales
Order Solitaria
Phylum Gastrotricha
Order Chaetonotida
Order Macrodasyida
Phylum Gnathostomulida
Order Bursovaginoidea
Order Filospermoidea
Phylum Micrognathozoa
Class Micrognathozoa
Order Limnognathida
Phylum Mollusca
Class Bivalvia
Subclass Autobranchia
Superorder Heteroconchia
Order Carditida
Order Lucinida
Order Myida
Order Pholadomyida
Order Trigoniida
Order Unionida
Order Veneroida
Superorder Pteriomorphia
Order Arcida
Order Limida
Order Mytilida
Order Ostreida
Order Pectinida
Order Pteriida
Subclass Protobranchia
Order Nuculanida
Order Nuculida
Order Solemyoida
Class Caudofoveata
Order Chaetodermatida
Class Cephalopoda
Subclass Coleoidea
Superorder Decabrachia
Order Sepiida
Order Sepiolida
Order Spirulida
Order Teuthida
Superorder Octobrachia
Order Octopoda
Order Vampyromorphida
Subclass Nautiloidea
Order Nautilida
(Continued)
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Class Gastropoda
Subclass Caenogastropoda
Order Littorinimorpha
Order Neogastropoda
Subclass Cocculiniformia (e.g., Cocculinidae)
Subclass Heterobranchia
Order Acochlidioidea
Order Anaspidea
Order Cephalaspidea
Order Gymnosomata
Order Hygrophila
Order Nudibranchia
Order Pleurobranchomorpha
Order Runcinacea
Order Sacoglossa
Order Stylommatophora
Order Systellommatophora
Order Thecosomata
Order Umbraculida
Subclass Neomphalina
Order N.N. (e.g., Neomphalidae)
Subclass Neritimorpha
Order Cycloneritimorpha
Subclass Patellogastropoda
Order N.N. (e.g., Patellidae)
Subclass Vetigastropoda
Order N.N. (e.g., Ataphridae)
Class Monoplacophora
Order Tryblidiida
Class Polyplacophora
Order Chitonida
Order Lepidopleurida
Class Scaphopoda
Order Dentaliida
Order Gadilida
Class Solenogastres
Superorder Aplotegmentaria
Order Cavibelonia
Order Sterrofustia
Superorder Pachytegmentaria
Order Neomeniamorpha
Order Pholidoskepia
Phylum Nemertea
Class Anopla
Order N.N. (e.g., Gorgonorhynchidae)
Class Enopla
Order Monostilifera
Order Polystilifera
Class Paleonemertea (e.g., Carinomidae)
Phylum Phoronida
(Continued)
48 / 60
Class N.N. (e.g., Phoronis)
Phylum Platyhelminthes
Subphylum Catenulidea
Order Catenulida
Subphylum Rhabditophora
Class Macrostomorpha
Order Haplopharyngida
Order Macrostomida
Class Neoophora
Subclass Eulecithophora
Infraclass Adiaphanida
Order Fecampiida
Order Prolecithophora
Order Tricladida
Infraclass Rhabdocoela
Order Dalytyphloplanida
Order Endoaxonemata
Order Kalyptorhynchia
Subclass Neodermata
Infraclass Cestoda
Order Amphilinidea
Order Bothriocephalidea
Order Caryophyllidea
Order Cyclophyllidea
Order Diphyllidea
Order Diphyllobothriidea
Order Gyrocotylidea
Order Lecanicephalidea
Order Litobothridea
Order Proteocephalidea
Order Pseudophyllidea
Order Rhinebothriidea
Order Spathebothriidea
Order Tetrabothriidea
Order Tetraphyllidea
Order Trypanorhyncha
Infraclass Monogenea
Order Capsalidea
Order Chimaericolidea
Order Dactylogyridea
Order Diclybothriidea
Order Gyrodactylidea
Order Mazocraeidea
Order Monocotylidea
Order Montchadskyellidea
Order Polystomatidea
Infraclass Trematoda
Order Aspidogastrida
Order Diplostomida
Order Plagiorchiida
(Continued)
49 / 60
Order Stichocotylida
Class Polycladidea
Order Lecithoepitheliata
Order Polycladida
Subclass Proseriatia
Order Proseriata
Phylum Rotifera
Class Eurotatoria
Subclass Bdelloidea (e.g., Adinetidae)
Subclass Monogonta
Order Collothecaceae
Order Flosculariaceae
Order Ploima
Class Pararotatoria
Order Seisonacea
Phylum Sipuncula
Class Phascolosomatidea
Order Aspidosiphoniformes
Order Phascolosomatiformes
Class Sipunculidea
Order Golngiiformes
Order Sipunculiformes
INFRAKINGDOM DEUTEROSTOMIA
Phylum Chordata
Subphylum Cephalochordata
Order Amphioxiformes
Subphylum Urochordata
Class Appendicularia
Order Copelata
Class Ascidiacea
Order Enterogona
Order Pleurogona
Class Thaliacea
Order Doliolida
Order Pyrosomida
Order Salpida
Subphylum Vertebrata [= Craniata]
Infraphylum Agnatha
Class Cephalaspidomorphi
Order Petromyzontiformes
Class Myxini
Order Myxiniformes
Infraphylum Gnathostomata
Superclass Actinopterygii
Class Chondrostei
Order Acipenseriformes
Class Cladistei
Order Polypteriformes
Class Holostei
Order Amiiformes
(Continued)
50 / 60
Order Lepisosteiformes
Class Teleostei
Order Acanthuriformes
Order Albuliformes
Order Alepocephaliformes
Order Anabantiformes
Order Anguilliformes
Order Argentiniformes
Order Ateleopodiformes
Order Atheriniformes
Order Aulopiformes
Order Batrachoidiformes
Order Beloniformes
Order Beryciformes
Order Blenniiformes
Order Carangiformes
Order Centrarchiformes
Order Characiformes
Order Cichliformes
Order Cirrhitiformes
Order Clupeiformes
Order Cypriniformes
Order Cyprinodontiformes
Order Elopiformes
Order Ephippiformes
Order Esociformes
Order Gadiformes
Order Galaxiiformes
Order Gobiiformes
Order Gonorynchiformes
Order Gymnotiformes
Order Hidontiformes
Order Holocentriformes
Order Istiophoriformes
Order Kurtiformes
Order Labriformes
Order Lampridiformes
Order Lepidogalaxiiformes
Order Lobotiformes
Order Lophiiformes
Order Mugiliformes
Order Myctophiformes
Order Notacanthiformes
Order Ophidiiformes
Order Osmeriformes
Order Osteoglossiformes
Order Pempheriformes
Order Perciformes
Order Percopsiformes
Order Pholidichthyiformes
(Continued)
51 / 60
Order Pleuronectiformes
Order Polymixiiformes
Order Salmoniformes
Order Scombriformes
Order Siluriformes
Order Spariformes
Order Stomiatiformes
Order Stylephoriformes
Order Synbranchiformes
Order Syngnathiformes
Order Terapontiformes
Order Tetraodontiformes
Order Uranoscopiformes
Order Zeiformes
Superclass Chondrichthyes
Class Elasmobranchii
Order Carcharhiniformes
Order Heterodontiformes
Order Hexanchiformes
Order Lamniformes
Order Myliobatiformes
Order Orectolobiformes
Order Pristiformes
Order Pristiophoriformes
Order Rajiformes
Order Squaliformes
Order Squatiniformes
Order Torpediniformes
Class Holocephali
Order Chimaeriformes
Superclass Sarcopterygii
Class Coelacanthi
Order Coelacanthiformes
Class Dipnoi
Order Ceratodontiformes
Order Lepidosirenoformes
Superclass Tetrapoda
Class Amphibia
Order Anura
Order Caudata
Order Gymnophiona
Class Mammalia
Subclass Prototheria
Order Monotremata
Subclass Theria
Infraclass Eutheria [= Placentalia]
Order Afrosoricida
Order Artiodactyla
Order Carnivora
Order Cetacea
(Continued)
52 / 60
Order Chiroptera
Order Cingulata
Order Dermoptera
Order Erinaceomorpha
Order Hyracoidea
Order Lagomorpha
Order Macroscelidea
Order Perissodactyla
Order Pholidota
Order Pilosa
Order Primates
Order Proboscidea
Order Rodentia
Order Scandentia
Order Sirenia
Order Soricomorpha
Order Tubulidentata
Infraclass Metatheria [= Marsupialia]
Order Dasyuromorphia
Order Didelphimorphia
Order Diprotodontia
Order Microbiotheria
Order Notoryctemorphia
Order Paucituberculata
Order Peramelemorphia
Class Reptilia
Subclass Aves
Infraclass Neognathae
Superorder Galloanseri
Order Anseriformes
Order Galliformes
Superorder Neoaves
Order Accipitriformes
Order Apodiformes
Order Bucerotiformes
Order Caprimulgiformes
Order Cariamiformes
Order Charadriiformes
Order Ciconiiformes
Order Coliiformes
Order Columbiformes
Order Coraciiformes
Order Cuculiformes
Order Eurypygiformes
Order Falconiformes
Order Gaviiformes
Order Gruiformes
Order Leptosomiformes
Order Mesitornithiformes
Order Musophagiformes
(Continued)
53 / 60
Order Opisthocomiformes
Order Otidiformes
Order Passeriformes
Order Pelecaniformes
Order Phaethontiformes
Order Phoenicopteriformes
Order Piciformes
Order Podicipediformes
Order Procellariiformes
Order Psittaciformes
Order Pteroclidiformes
Order Sphenisciformes
Order Strigiformes
Order Suliformes
Order Trogoniformes
Infraclass Paleognathae
Order Apterygiformes
Order Casuariiformes
Order Rheiformes
Order Struthioniformes
Order Tinamiformes
Subclass Crocodylomorpha
Order Crocodylia
Subclass Rhynchocephalia
Order Sphenodontida
Subclass Squamata
Order Anguimorpha
Order Gekkota
Order Inguania
Order Lacertoidea
Order Scincoidea
Order Serpentes
Subclass Testudinata
Order Testudines
Phylum Echinodermata
Subphylum Asterozoa
Class Asteroidea
Order Brisingida
Order Forcipulatida
Order Notomyotida
Order Paxillosida
Order Peripoda
Order Spinulosida
Order Valvatida
Order Velatida
Class Ophiuroidea
Order Euryalida
Order Ophiurida
Subphylum Crinozoa
Class Crinoidea
(Continued)
54 / 60
Order Comatulida
Order Cyrtocrinida
Order Hyocrinida
Order Isocrinida
Subphylum Echinozoa
Class Echinoidea
Subclass Cidaroidea
Order Cidaroida
Subclass Euechinoidea
Infraclass N.N.
Order Echinothurioida
Infraclass Acroechinoidea
Order Aspidodiadematoida
Order Diadematoida
Order Micropygoida
Order Pedinoida
Infraclass Carinacea
Order Arbacioida
Order Camarodonta
Order Salenioida
Order Stomopneustoida
Infraclass Irregularia
Order Cassiduloida
Order Clypeasteroida
Order Echinolampadoida
Order Holasteroida
Order Spatangoida
Class Holothuroidea
Order N.N. (Thyone)
Order Apodida
Order Aspidochirotida
Order Dendrochirotida
Order Elasipodida
Order Molpadida
Phylum Hemichordata
Class Enteropneusta (e.g., Harrimaniidae)
Class Pterobranchia
Subclass Cephalodiscida (Cephalodiscus)
Subclass Graptolithina
Order Rhabdopleurida
Phylum Xenacoelomorpha
Subphylum Acoelomorpha
Class Acoela (e.g., Diopisthoporidae)
Class Nemertodermatida (e.g., Nemertodermatidae)
Subphylum Xenoturbellida
Class N.N. (Xenoturbellidae)
Names below rank of infrakingdom are arranged alphabetically within each parent rank, except for taxa that are not named (N.N.). Brackets indicate
synonyms. Quoted names are not validly published but in common use.
doi:10.1371/journal.pone.0119248.t002
55 / 60
the many taxa therein and provides for easier import and manipulation of data by
information systems.
Supporting Information
S1 Appendix. List of sources consulted for proposed higher level classification of all living
organisms.
(PDF)
S1 Table. Proposed hierarchical classification from superkingdom to order.
(XLSX)
Acknowledgments
We thank those on the expert panel who are not authors of this paper for their valuable contributions earlier in this process. In addition, we are very grateful to David Nicolson for his help
in reviewing and improving the many drafts of the classification, Brian Tindall and Peter Stevens for their helpful review of the draft manuscript, and the three PLoS reviewers for their useful comments. We also thank Gloria Ruggiero for her excellent editorial work on
the manuscript.
Author Contributions
Conceived and designed the experiments: MR DG NB TB RB TC-S MG PK TO. Analyzed the
data: MR DG NB TB RB TC-S MG PK TO. Wrote the paper: MR DG NB TB RB TC-S MG PK
TO.
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A Higher Level Classification of All Living

HCMR - Hellenic Centre for Marine Research,

Musum National d'Histoire Naturelle

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The University of Arizona

National University of Ireland, Galway

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Available from: Thomas M. Orrell

A Higher Level Classification of All Living

Data Availability Statement: All relevant data are

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

morphological; molecular: genes, metagenome, and metabolome; etho-ecological). There is

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

Results and Discussion

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

included a subkingdom Archaea; Cavalier-Smith also treated Archaebacteria and Eubacteria as

Protozoa and Chromista

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

Table 2. Proposed hierarchical classification from superkingdom to order.

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms

PLOS ONE | DOI:10.1371/journal.pone.0119248 April 29, 2015

A Higher Level Classification of All Living Organisms